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Co-evolution of Humans and Canids
An Alternative View of Dog Domestication: Homo Homini Lupus?
Why did our ancestors Dogs and wolves are part of the rich palette of predatame and domesticate tors and scavengers that co-evolved with herding unwolves, of all creatures, gulates about 10 Ma BP (million years before and turn them into dogs present). During the Ice Age, the gray wolf, Canis luto become man’s best pus, became the top predator of Eurasia. Able to keep friend? Is man dog’s best pace with herds of migratory ungulates wolves befriend, as Mark DERR came the first mammalian “pastoralists”. (DERR 1997) once dared to Apes evolved as a small cluster of inconspicuous treedwelling and fruit-eating primates. Our own species ask? Well, not according separated from chimpanzee-like ancestors in Africa to Stephen BUDIANSKY’s around 6 Ma BP and– apparently in the wider context assertions in a new book, of the global climate changes of the Ice Age–walked as The Truth About Dogs true humans (Homo erectus) into the open savanna. (BUDIANSKY 2000). He Thus an agile tree climber transformed into a swift, claims that dogs are scavcursorial running ape, with the potential for adopting engers at heart “For all the migratory life style that had become essential for the myth and tales of Co-Evolution of the inhabitants of the savanna and steppe. In the abdog’s service to man, only Cooperation: An sence of fruit trees, early humans turned into omnivoa small fraction of dogs rous gatherers and scavengers. They moved into the Alternative to living off human society steppe of Eurasia and became skilled hunters. Domestication? today earn their keep… Sometime during the last Ice Age, our ancestors The human species has the overwhelming majorteamed up with pastoralist wolves. First, presumably, reﬁned sociality to a deity of dogs were freeloadsome humans adopted the wolves’ life style as herd gree of complexity that ers” (p6), even though “in followers and herders of reindeer and other hoofed anremains unmatched in my role as brutally objecimals. Wolves and humans had found their match. our world. The human tive observer, I do love We propose that first contacts between wolves and capacity for cooperation dogs” (p9). As scientists, humans were truly mutual, and that the subsequent starts at birth: even we are skeptical about a changes in both wolves and humans are understood though a human mother successful journalist who best as co-evolution. is able to give birth and claims to have found “the take care and full respontruth”, especially in such Key words sibility for the newborn, a complex web of probChimpanzee, co-evolution, cooperation, dog, Eurasia, traditionally three lems, connections, obserhuman, hunter, Ice Age, pastoralist, Pleistocene, sowomen cooperate in the vations, opinions. And, cial, pack, wolf. birth… the mother, the his confession of his mother’s mother and the “love” for dogs has a midwife. Growing up is sheltered by complex helper strange contradictory ring. systems, staffed by siblings, babysitters, and teachKonrad LORENZ once stated: “Of all creatures the one nearest to man in the ﬁneness of its perceptions and ers far beyond puberty, and group living within the in its capacity to render true friendship is a bitch.” family is well supplemented by socialization among
Evolution and Cognition
(LORENZ 1954, p85). Isn’t it strange that, our being such an intelligent primate, we didn’t domesticate chimpanzees as companions instead? Why did we choose wolves even though they are strong enough to maim or kill us? We do not claim to know “The Truth” but we offer in this paper a different view, with emphasis on companionship rather than human superiority.
2003, Vol. 9, No. 1
The high morality we claim as achievement of our species. and even ethics (SCHLEIDT 1998). the African Cape hunting dog and the Asian dhole. not only in well coordinated drives in the context of attacking prey. we ﬁnd ourselves in a strange conﬂict. The closest approximation to human morality we can ﬁnd in nature is that of the gray wolf. and that various subsequent changes in both wolves and humans must be considered as a process of co-evolution. This is especially odd in view of the bad reputation wolves have in our folklore. or the persecution of a competitor. we propose that initial contacts between wolves and humans were truly mutual.. from kindergarten. Schleidt/Michael D. habits.1 There a new species emerged and dominated Europe and the Near East longer than any other ❘ 58 ❘ 2003. and became ﬁerce hunters during the grueling conditions of the Ice Age. but the gravity of the problem became apparent when the genetic relationship between wolves and dogs was elucidated by Robert WAYNE and Carles VILÀ (VILÀ et al. The life of chimpanzees. Even in their maternal behavior warmth and affection are apparently reduced to nursing and an occasional comforting hug. In Thomas HOBBES’ own words: „To speak impartially. Such a long common history of dogs and modern humans begs the question as to the dog’s part in the endeavor of humans to take control of the world. 1 . kinship. 1997). and led to the formulation of a hypothetical “lupiﬁcation” of human behavior. predating human sociality and cooperativeness by millions of years. and ending in funeral rites.000 years before present). etc. as proposed for hoofed animals and fowl. FOX 1975. carrying items too heavy for any one individual. both sayings are very true. however. even though for a rancher who leaves his livestock unsupervised and unprotected by a good shepherd. and meet our maker who made Adam in His likeness. 1980. rather.. Our own paper does not aspire to be a comprehensive review of the hypothetical co-evolution of cooperation between dogs and humans. our understanding of wolves has changed considerably. and. and that Man to Man is an arrant Wolfe. Canis lupus. MECH/BOITANI 2003). humans created a god in their likeness who is their alter ego. especially their sociality. Wolves’ ability to cooperate in a variety of situations. That Man to Man is a kind of God. Or. provisioning not only their own young but also other pack members. in the absence of any historical evidence turn to our closest relatives. work. 9. Erhard OESER has pursued this lead and traced the contribution of dogs in the “humanization of the ape” in a wider context of human culture (OESER 2001. similar forms of cooperation are observed in two other closely related canids.Wolfgang M. is rivaled only by that of human societies. HOMO HOMINI LUPUS. cooperation among group members is limited to occasional hunting episodes. In addition. Therefore it is reasonable to assume that canid sociality Evolution and Cognition and cooperativeness are old traits in terms of evolution. baby sitting. Homo sp. we can give a new and very different meaning to HOMO HOMINI LUPUS: “Man to Man is—or at least should be—a kind Wolfe. and accepted by many as the ultimate judge of their moral behavior. opening the possibility that the split between wolves and dogs may date back as far as 100 to 135 ka BP (135. Primates and Canids: A Current View Around 6 Ma BP humankind separated from chimpanzee-like tree dwelling and fruit-eating ancestors in Africa and moved as true human hunters and gatherers. Thus. the chimpanzees. as revealed by the pioneering work of Jane GOODALL and others (GOODALL 1986. however. DE WAAL 1997) appears as a frightful caricature of human egoism. an “arrant Wolfe” is to this day a formidable threat (MECH 1970. and our newspapers are full of stories that reﬂect more chimpanzee than human ethics. through school. into the open savanna. is a very thin veneer on the old ape. Instead of perpetuating our traditional attitude that our ”domesticated animals” are intentional creations of human ingenuity. The impact of wolves’ ethics on our own may well equal or even exceed that of our effect on wolves’ changes in their becoming dogs in terms of their general appearance or speciﬁc behavioral traits. in press). The ﬁrst insight we get from chimpanzee society is: “We have come a long way”. When we try looking back at the biological foundations of our moral behavior in a distant past. can be found in ZEUNER’s pioneering work (ZEUNER 1963). but attempts merely to provide more details und pursue several collateral ideas emanating from the original proposal (SCHLEIDT 1998). Shalter peers. always aimed for one’s own advantage and executed with MACHIAVELLIAN shrewdness. “(HOBBES 1651). conquered the plains of Eurasia. Vol. No. But our communal life does not end there: We strive to join our ancestors in heaven. as expressed in the famous phrase.” This shift in our attitude toward wolves opens a new vista as to the origin of dogs. The earliest suggestions that dogs do not ﬁt the conventional paradigms of domestication. Since HOBBES’ time.
the legendary African Eve. and around 10 Ma BP.000. Back to serious science: The canids. Africa. wrote in a letter to the ﬁrst author: “Dogs have been domesticated for a very long time.000. coexisting and competing with the big cats and hyenas: the wolves. So the story goes. Then.). killed all the reindeer. Jane GOODALL.000 Aelurodon stirtoni Hesperocyoninae 100.000 recent 10 Miocene 20 Caninae 10. and the simplistic view of Pleistocene as an epoch of inhospitable cold climate in circumpolar regions. The Genus Canis apparently evolved in Asia. Thus. hominid: the clumsy. mammoths and Neanderthals. He took to the daughters of African Eve who ﬂourished in icy Eurasia. they started to cross into Asia. but about what we may poetically associate with kindness of heart.000 Tertiary 60 Paleocene The cast of human and canine evolution With fruit trees and reindeer as parts of the stage set Figure 1: Cladogram indicating the phyletic relationship between Canis lupus and selected species and divisions of Canidae within their temporal and geographic ranges (after TEDFORD/TAYLOR/WANG 1995. man tamed wolves. latrans Ma 0 Plei. and by artiﬁcial selection created the multitude of dog breeds we know today. Figure 2: A schematic summary of the current teaching of human and canid evolution. Here we are not talking about intelligence. coyotes (the latter all members of the genus Canis). as a species distinct from the various jackals. etc. WAYNE/ VILÀ 2001. invited to comment on K. aur. MCKENNA/BELL 1997. Or as an alternative hypothesis. which were scavenging among the rich refuse of human camp sites. and the aberrant “wild dogs”. Vol. around 150 ka BP—based on evidence from mtDNA—a superior woman. den together. A multitude of swift canid predators evolved. emerged from the forest and were originally about the size of a fox when grasslands opened and herds of grazing ungulates. too. Plio. the circumpolar big “wolf”. 3 They. They hunt together. Red foxCuon like Canids North America Red foxlike Canids Urocyon Africa Eurasia North America Canis lupus C.000 Pleistocene 30 Oligocene 1. 9. they became part of the rich palette of canid predators and scavengers. Her daughters moved into icy Europe and. began to dominate the open plains. had emerged. They have descended from wolves who were pack animals. have their roots in North America. 1 .000 40 Eocene 50 10. the ﬁrst man who really deserved the name “Homo sapiens”. and Canis lupus. thanks to their rich subcutaneous fat and superior intelligence.2 Meanwhile. by pretending to be their obedient servants and hunting companions. Europe. and back into North America. Primates and Canids: A Different View There is something in the bond among wolves and between dogs and humans that goes beyond that between us and our closest primate relatives. were able to outcompete Neanderthal women. LORENZ’s statement quoted above. around 80 ka BP—based on evidence from nuclear DNA—the legendary African Adam emerged.Co-evolution of Humans and Canids Africa C. No. appeared about one million years BP. and did beautiful art-work in the caves of Spain and France. Note the logarithmic time scale. jackals. Borophaginae Years before present 1. also known as (wild) “dogs”. They survive as a result of teamwork. the African Cape hunting dog. scavenging wolves took the initiative and conned the afﬂuent hunting and gathering humans into sharing their plenty. the chimpanzees. Evolution and Cognition and the Asian dhole. simensis Lycaon Eurasia Canis lupus C. raise pups to- ❘ 59 ❘ 2003. At the end of the ice age. but successful Neanderthal man. notably horses and antelopes.
Schleidt/Michael D. community. but the desire to help. but able to squeeze by on the diet and life style of a fox: hunting mice and picking berries. And even between family members. but in denning together and raising pups together. The ubiquity of the gray wolf is apparently due to its rich behavioral repertoire and the ability to adapt its life style opportunistically to local and temporal conditions: most successfully as a pack hunter of midsize ungulates. The E PLURIBUS UNUM of the pack goes far beyond what makes UNUM. As in the social insects (bees and ants). including loyalty. ness and MACHIAVELLIAN reasoning get in the way of our behaving communally. when a stranger or even a close kin fails to comply. and in India it was sympatric with the dhole. and agree on a common goal. In fact. they are stronger than a single person. It is not related to intelligence. the dhole of India (Cuon alpinus) and. to be the highest achievement of humanity. you see the love between mother and offspring. In some areas the gray wolf coexisted with less social members of the genus. but maybe even more so by the wild dog of Africa (Lycaon pictus). even for intelligent human beings. we admire the age-old saying. It is practiced to this very day by some of their descendents and honed to perfection by members of the pack-hunting canid species: notably the gray wolf. But also. As a rule. or even foxes. 1 . out of a herd or a gang of selﬁsh ﬁghters. at least in theory. pack members can also accept strangers. and nation. and the bonds between siblings. to be obedient. 3) (HARRINGTON/PAQUET 1982). nuzzling each other. Several species of canids show these special traits. or force them to comply.Wolfgang M. however: the members of the group must cooperate. there are indications that such humaneness. Strangely. to gain our approval. the gray wolf. on rare occasion. anatomical feature. disputes often rise that may even lead to ﬁghts… even after hundreds of years of selective breeding. Self interest ﬁrst. There is a catch. which many admire and hold. it can be attacked. We not only value a dog’s intelligence. a hero or saint can overcome temptations of selﬁshness. as they have lost access to big herds of ungulates and now tend more toward a lifestyle similar to their “minor brothers:” coyotes. That is not as easy as it sounds. Shalter gether. where the hive is only one of the units of selection (MORITZ 1993)5. and loyalty. was invented millions of years ago by early canids. if two or more persons can agree to cooperate. the bigger the group.6 What is it that makes the ancient pack social system so successful? Well. Let others take the risk and reap the gain for oneself and one’s kin. In theory. we praise it as the highest expression of humaneness when. jackals. some of today’s wolves may well be less social than their ancestors. it would be hard if not impossible to produce a chimpanzee who could live with humans and have anything like such a good relationship as we have with our dogs. They are boisterous and volatile in the wild. and if there is a little surplus: practice nepotism. one. Sweet it is to die for one’s country. communicate. If you watch wild chimps. many peoples. also called the Cape hunting dog (Lycaon pictus). the easier it can subdue a single person or a few. Jane GOODALL’s eloquent comparison is based not only on her own personal experience with dogs 4 and many years of living with chimpanzees (GOODALL 1986). not only during the hunt. Many people. wagging their tails in greeting. the African brother of our gray wolf. it is not a single life history trait. We preach love thy neighbor and ask our brave boys in uniform to be prepared to sacriﬁce their lives for the sake of their families. Equally. The old primate trait of selﬁshEvolution and Cognition ❘ 60 ❘ 2003. 9. driven off. licking and protecting the pups. They do superb teamwork. That is the basis of majority-ruled democracy. They are not pack animals. a unit . one nation. Chimpanzees are individualists. but its warm affection. or type of physiology or E Pluribus Unum From many. the pack became one of many systems upon which natural selection acts. you see all the characteristics we love in dogs. Although the core of the pack is usually one extended family. playfulness. of course. They are always on the lookout for opportunities to get the better of each other. the bush dog of South America (Spethos venaticus). and several other really outstanding behavior patterns related to harmonious life in a pack. Vol. to a lesser extent. but also less well known to the general public on her familiarity with and research on the wild dog. or even killed. This ancient social order has been helpful in the domestication of the dog. Humane Canids Among all the canids one species became the most successful mammalian predator ever: Canis lupus. No.” (GOODALL 1997). If you watch wolves within a pack. It roamed over all the northern hemisphere north of 15˚N (Fig. Other relationships tend to be opportunistic.
and a skeleton of this species on display in the Museum of Natural History of the Smithsonian Institution in Washington. In wolves each pack member can accept greater risks when attacking. and a low ranking pack member may have to wait. do they incur high risk of getting seriously injured. and for Grimm’s fairy tale bad wolf swallowing grandmother and Little Red Ridinghood). for example. they are able to run as a single group. or when they get home to the den. Wolﬁng down prey is apparently an ancient canid trait: Around 11 Ma BP a wolf-type canid ( Strobodon stirtoni) roamed Nebraska. D. The attacks on prey by lions. as a rule. but which individual. The second phase of feeding starts when the wolves have reached a cozy place for a rest some distance from the kill. sharks. but among individuals bonded as mated pairs or by lasting friendships among individuals of the same gender. only one pair breeds. or for two dogs to race at full speed while holding onto a stick of wood. as. but compared to other predators there is little overt competition among pack members. a female). they can. No. there is growling and snarling. will get “the lion’s share”. Typically predators. is the central feature of canid pack living. tigers. they are low-risk performances by smooth butchers. They then regurgitate the large chunks. All is tuned to swallow as much as possible as fast as possible (which is the basis for the story of Fenris-Wolf gulping down Odin. when going for the kill. It is a whole array of speciﬁc adaptations which make communal life possible. behavior. When wolves feed on a kill. where the individual that makes the kill and can maintain possession of the carcass. apparently well aware not only that another pack member is running where. and vultures arrive. which allows pack members to make maximal use of each kill and to leave little for others. Social packforming canids are essentially monogamous. when injured.Co-evolution of Humans and Canids Gray Wolf: current distribution Gray Wolf: now extirpated Figure 3: Grey wolf (Canis lupus) (after HARRINGTON/PAQUET 1982). the needy Evolution and Cognition will be fed by the other pack members. sharing ❘ 61 ❘ 2003. When canids hunt as a pack. This awareness makes it possible for a team of dogs to pull a sled and run for hours without changing places. where the strong tend to take from the meek). leg bones articulated and neatly folded-up in the area where once the canid’s stomach had been. in conspeciﬁc ﬁghting over a limited resource (e. however. Even siblings and friends share food and affection (unlike in chimps. or take it over by force.8 reveals an amazing story: in the region of the ribcage one can see quantities of broken and etched bone representing at least two individuals of small antelope.g. avoid the risk of disabling injury that would prevent them from hunting. lions. hyenas. of course. act much more as an integrated system than any group of chimps or lions. hyenas. but all members share food and parental care generously. 1 . there is usually not much left. Only when they turn on each other. Even though there may be several sexually mature adults in a pack. 9. In reality. By the time jackals. Wolﬁng down prey is but the ﬁrst phase of feeding. and the like conjure up images of bravery and fury. The long-legged social canids are not only fast and long-distance runners. Vol. because.7 This cooperation and risk sharing not only among close relatives. because of their focused attention and close cooperation. tigers.C.
very much like you can determine what a dog listens to by observing its ears provided they are pointed like those of wild canids (SHALTER/FENTRESS/ YOUNG 1977). Shalter with those that did not participate in the hunt. jumping. a hare—prey up to about one’s own body size—eating a few berries and a couple of mushrooms along the way. where serious threat by a high ranking member is rare compared to the gracious acceptance of a lower ranking individual’s signs of compliance or even submission. Thus. we are easily led to believe that our ancestors had nothing better to do than to leave their beastly existence ❘ 62 ❘ 2003. There are other behavioral features of importance for the canid pack algorithm. The pack hunter’s social awareness is equally amazing. sheep. the social systems of canids and their hunting strategies can be considered a continuum from fox to wolf. But. just about every wolf was shot on sight? What about the lone wolf? Human wolf eradication programs go for the conspicuous individuals. they can be so overwhelmed with joy that they bounce around and prance off together as if they were thinking: Let us prey! Thus. Lupiﬁcation of Canids When we talk about our own primate descent. Vol. and a pack raiding a sheep shed or bringing down a cow in a pasture is much more conspicuous than a single wolf that is forced to adopt the lifestyle of a fox.g. they keep pieces for themselves. The pack at the den can process its loot in peace and spend time resting and digesting. for the past several centuries. depending on who is going where. and who is doing what. multi-species packs: humans. e. if they prefer. The African hunting dog and the dhole of India may be even a bit further out to the extreme of pack life. This awareness is an essential feature of both: enabling dogs to ﬁt so well into our human social fabric and enabling the pack-hunting wolves to lead a communal life: moving and hunting together. The same applies to smart dogs: do not believe that opening the fridge or reaching for a can opener triggers a mere Pavlovian conditioned reﬂex in your dog. For example. such as a leg of a prey. with “real growls”. when an individual buries its leftovers. in spite of all the generous sharing of food. listening and snifﬁng. for example. and start to dig only when nobody is in sight.g. Contrary to the popular belief that canids are specialized in snifﬁng and have limited eyesight. they constantly watch each other. horses living in harmony. Finally. consider such doggish behavior—accommodating rather than ﬁghting— as cowardly. enforcement of established rank can be a low-key affair. Occasionally ﬁghts break out. Like wolves. in hyenas. we may ask: What about wolves at very low population densities as in most of Europe. and very different from the ﬁghting over a kill. a life which only few. What had been carried communally. goats. MACHIAVELLIANS. the wolves surviving at very low densities may well have lost their social competence. with the coyote and jackal somewhere in between. and chasing down whatever one can grab: a frog. if any. where.. a mouse. it appears they use the basic bag of canid tricks that foxes are known for: wandering around low proﬁle. Their social skills are practiced only toward their mates and young. it can behave very secretively. and even close relatives can be killed. etc. the special success of the gray wolf may well be based on this species’ potential to live well like a dhole and still survive when forced to live like a coyote. too large to swallow. Schleidt/Michael D. each member of the pack knows not only who is who but also who is where and who is doing what. This is what in dogs makes it easy for wolﬁsh families to form mixed. both in their genetically endowed propensities and in their social pack-living skills. but only rarely does one observe strict rank orders and real macho behavior on the part of dominant pack members. then stalking. looking. the movement of the eyebrow (the light spot above a Doberman’s eye or in the face mask of a husky) will tell you what it attends to. Yet it is precisely what keeps canid pack members from incessant quarreling. cats. about the hominization of Australopithecines. 1 . or from playing macho the way chimps or some humans cannot do without. as. If you question our claim and are hung up on your belief in Pavlovian conditioning. dogs are also very much aware of who is who. and carefully go over what they brought home in their stomach shopping bags. dogs. but actually quite playfully. You will quickly realize that the conditioned reﬂex explanation for a dog’s awareness is a gross simpliﬁcation. and pulled apart in a “tug of war”.Wolfgang M. who is where. sharing. you will see its eyes move. 9. especially the pups and their babysitter. is cut down to size. of our wolves can indulge in today. when two lone wolves meet. the way hyenas do. No. e. If you watch it carefully. When hunting. dealing with hierEvolution and Cognition archy with minimal bloodshed. train your dog to bring speciﬁc items (toys you assign names to) and then give each to a different member of your family or deposit them at different locations. Instead. And if you cannot see its eyes. however. And. a rabbit.
Only during the last few thousand years did humans propel themselves in mass to the top of the food pyramid. under the heading Humane Canids: what made the ancient pack social system so successful was not a single life history trait. As these herbivores responded by joining together in large herds. however. The predators latched onto this new and evolving ecosystem. 9. Canine Humans? So. we ﬁnd comfort in our cherished belief to be fruitful. When the land bridge united North America with the Asian continent in the area we call the Bering Strait. Weichsel in northern Europe and Würm in the European Alps) peaked only 18. others still behave like chimps or lions. etc. had a climate in Europe warmer than today. especially the image of northern Eurasia and America being covered by one gigantic ice shield that only temporarily gave way to a little green during the interglacials. developing communal defense behaviors. when the pack hunters moved into Asia and Africa. with a degree of omnivorousness. climate and ecology. and a shewolf as a member of a wolf pack? Let us look. the grueling cold of the last phase of the Wisconsin (in America. grasses started to cover large areas of the earth. point by point. Fig. and over every living thing that moveth upon the earth. with many niches for canine companions. and an evolutionary arms race started between grazers and grasses. some of its ancient predators specialized into fox-like carnivores. 4) are most important variables in the evolutionary puzzle of the genus Canis. if they failed to kill them. They grew larger and stronger. but were at the mercy of the new pack-hunting ancestors of wolves. A blind person may depend on a seeing-eye dog.. In other words. the Ipswichian of England. Vol. The horses. When that continent became disconnected from the other land masses. to the size of coyotes and larger. villages to cities. and subdue it… to have dominion over the ﬁsh of the sea. Asia. multiply. instead of seeing ourselves as part of the complex system of nature. early horses and other ungulates changed from leaf eaters to grazers. nations. Not only did they compete for the resources formerly controlled by lions and tigers. what would they ﬁnd? How would wolves view the lupiﬁcation of their canid ancestors? As Africa is today considered to be the cradle of mankind. about 135 ka BP until about 70 ka BP. fox-like early canids had to catch up with their prey. When.000 years ago. Many humans behave the canine way. snifﬁng at the old bones of their dead and the bones left of their meals. did exceedingly well in the open grassy plains of Asia and Africa. the smart. at least they killed their cubs. which had adapted well to predation by felids. and. possibly driven by changes in global climate. and dholes. with hippopotamus wallowing in the Thames and Mediterranean vegetation ﬂourishing in the valleys of the Austrian Alps.Co-evolution of Humans and Canids behind and let those not worthy of becoming “humans” die out (Neanderthals. even among siblings and friends. and the canids thrived on the native herbivores. Thus. as a member of our household. anatomical feature. as fast runners. roughly as monogamously as most members of wolf packs. they joined or even replaced the big cats at the top of the food pyramid. or matter of physiology or behavior. insectivores. slower grazers and especially on their young. Humans. On the other hand. horses and canids poured into Asia. the ﬂeas) is traceable to North America. we continue to pretend to be the very crown of creation. and insects. canids “invented” long legs for high speed running. what is the difference between a beloved golden retriever bitch. but a whole array of speciﬁc adaptations that make communal action possible. the prerequisite for hunting in a fast moving pack. and North America. at what we said above. the origin of canids (as well as their old parasites. family bands. The Ice Age as the geological epoch and the “Mammoth Steppe” as the biogeographical substrate (GUTHRIE 1990. As a counter strategy to the herding of ungulates around 10 million BP. Our spectrum includes the solitary existence of saintly hermits. live more or less monogamously. of previous teachings were wrong. displacing the canid pack hunters. 1 . or the like). and over the fowl of the air. We are still far from a consensus about the causation and dynamics of Pleistocene geology. and outgrowing their predators in size. Our understanding of these variables has changed dramatically in our lifetime. In the human species we ﬁnd a very wide range of family structures and a great variety of even more complex (“non-family”) social systems. We share parental care generously. not necessarily humanely. No. more recently. at least in Western cultures. if not most. but it has become obvious that many. The last interglacial. hunting dogs. bushmen. If wolves could dig up the dens of their ancestors in Europe. a family dog ❘ 63 ❘ 2003. preying on the smaller. replenish the earth. the genus Homo as well. presumably living on small rodents. In spite of accepting the new creed of Darwinian natural selection . but they Evolution and Cognition even attacked them and.
hunting behavior by dogs is discouraged and subject to artiﬁcial selection: where local leash laws are not respected. most dogs’ legs compared to those of wolves or working dogs are not suited for running and are even somewhat crippled.S. it stays down. They let it sniff ﬁre hydrants. Bernard. Thus. On the other hand. when varying parts of grassland were replaced by forests (after GUTHRIE 1990). ancient mode: e.5 million years and had been more extensive during glacial episodes and less so during interglacials.. No. A dog knows not only who is who but also who is where and who is doing what. face an uncertain future in a pound. many dogs prefer to lie on a windowsill and watch the world go by.. etc. and the strange air when riding along in a car with its nose stuck out the window. trees. Joining dogs in communal hunting behavior was long reserved for pharaohs. This speciﬁc set of skills is a prerequisite for a dog’s ability to know and recognize many different people. The only exceptions are a few breeds of so-called hunting dogs. or. as have most city dwellers. Once it’s down. chewing toys. Not only dachshunds. Ulysses returning home in rags after many years away and being recognized and ❘ 64 ❘ 2003. work/guard dogs (Dobermann. Shalter Mammoth steppe Glaciation Figure 4: The “Mammoth Steppe” in the wider context of Pleistocene glaciation. bulldogs. fox terriers. etc. Vol. Many a German shepherd champion is no match for a husky or greyhound on a long-distance run. we remind you once again: contrary to the popular belief that canids are specialized in snifﬁng and have limited eyesight. at the very least. 1 . and the upper class. But otherwise. there is no need for him to regurgitate and look it over again. but they walk their dog around the block several times a day. as sport for a few rich. and a Dalmatian may not only serve as a ﬁre department’s helper in situations where humans are bound to fail. so long as they “work” under close human supervision. shepherds.). Schleidt/Michael D. Although Fido still wolfs down his well-processed boneless meal. piece of hide. strays are shot by game wardens or park police. or a specially processed bone. as sport for the common man. (hounds. and the sporting group (beagles. in the U. Feeding on the kill? Well. they are constantly watching each other. who no longer rely on their feet but on cars and elevators. Humans run neither as fast nor with as much endurance as wolves.). terriers). having lost their wolﬁsh striving for long-distance locomotion. and only exceptionally in the original. biking. golden retrievers. Nowadays it is pretty much restricted to traditional Old English fox hunts with a pack of hounds. what is in the food dish rarely involves much effort or opportunity to share or compete in a one-dog household. but also a beloved mascot. as well as squirrels. The extent of the steppe varied over the last 2.Wolfgang M. can be a great asset for a youngster. To ensure our dogs’ good teeth we buy special tartar-control treats. royalty. dried pig’s ear or ox penis! The dog’s social awareness is one of its greatest assets. thanks to the grinding action of the dogfood industry. some still hunt with a pack of hounds—black bears and mountain lions.g. St. But today’s stereotypes are the American hunter with riﬂe rack in Evolution and Cognition his pickup truck and his retriever next to him. to greyhound racing. and Yorkshires act as if hobbled by their anatomy. Maltese. 9. or hiking. or take it along jogging. Therefore. or the German forester with pipe and dachshund.
humans are apparently the only primates to make use of caves. But where do we draw the line between ancestral species? Could we interbreed with Neanderthals? In canids. in the context of human activity are dated about 14. the mother common to all modern humans. And although dogs perform no obvious farewell ceremony. or geneticist. 1 . the red wolf. many show signs of sadness and depression after a beloved friend or any member of its family (its pack) has left. paleontologist. provided there is a certain level of social competence among the human pack members. lupus. some scientists believe that this hypothetical Eve lived 160 ka BP in Africa. sheep. based on the analysis of mtDNA (mitochondrial DNA) from various canids. however. WAYNE’s data indicate that dogs are related closest to wolves found today between France and western Russia. and not to those in the Near East. and cattle (BENECKE 1995. C. 1999) at the laboratory of Robert WAYNE at UCLA. Asia.g. 9. based on the same reasoning that traces our Eve and Adam back to Africa. were our own ancestors back then. A word of caution. a foreshortening and rounding of the skull. but it will take a while before we come close to reconciling the many contradictory “facts”. and genetic distance can give us some idea of the elapsed time since two forms separated. Genetic differences between wild and domesticated forms can be estimated in various ways. Are such domesticated forms now members of a different new species? Let the professional taxonomists ﬁght that out. although supported by considerably larger data sets than many estimates of our own ancestry. ethologist. however. C. and many others had assumed previously). e. geneticists have developed a number of tests as measures for the degree of relatedness by comparing differences in the DNA of individuals and populations. and faster sexual maturation. all species of the genus Canis can interbreed. Based on such studies in humans. and their association with dogs predates the construction of permanent houses by thousands of years. This study. Once the ranges of variability of the assumed wild ancestors and those of the ancestors of our domesticated forms no longer overlap substantially. In recent years. latrans (WAYNE/JENKS 1991). for example. (1997.9 Domestication: Who Domesticated Whom? The oldest remains of dogs. we have a different problem: apparently. It can get tricky. depending on whether you ask a morphologist. long before any trace of domesticated goats. was performed by Carles VILÀ et al.. we assume that the exchange of genetic material between those two populations was reduced or interrupted by human interference and control. most dogs have no reason to even attempt to take over to become top dog and run the family.Co-evolution of Humans and Canids greeted by his old dog has become the archetypical experience that repeats itself daily at many airports and train stations. No. Vol.000 years BP. SERPELL 1995). less “wild” than the wolves they associated with? While canids are known to dig their own dens. SABLIN/KHLOPACHEV 2002). and the coyote. niger. that is. and North America). has revealed itself as a stable hybrid between the gray wolf.” But. The most Evolution and Cognition common criterion for belonging to one species is interbreeding and having fertile offspring. Is it not absurd to talk about the “domestication” of dogs by humans who had not yet any permanent domiciles (“domus”)? What are the signs of domestication in the archeological record? In mammals it is usually a reduction in overall size. where many of our domesticated animals and plants are assumed to have originated. There is less agreement concerning the ﬁrst Homo sapiens. mtDNA from 67 breeds of dog showed a high degree of similarity to that of wolves (as represented by 27 wolf populations from Europe. it has been assumed that the common ancestors of chimpanzees and early man split about 6 Ma BP. even more controversial. ❘ 65 ❘ 2003. even over hundreds of years (THOMAS 1993). long before they built permanent houses for themselves. Thus. of a canid distinctly different from wolves. nearly TEN times further back than indicated by any bones or any paleontological evidence found thus far! And. what do we mean by “domesticated”? In a most general sense: “no longer in its wild or natural state.10 New results. indicate that the split between the ancestors of wolves and jackals occurred about 1 Ma BP. clearly supporting the hypothesis that dogs are descendent from wolves (not from the golden jackal or other canids. and some of such dens may have been used by many generations. But the most spectacular of WAYNE’s results is that the ﬁrst split between dogs and wolves dates back roughly 135 ka BP. as Darwin. A good family dog is well integrated into the social hierarchy and. and our hypothetical Adam lived there 70 ka BP. C. particular observations of the fossil and genetic evidence. there is little argument that dogs are the oldest domesticated animal (ZEUNER 1963.
tubers. and reindeer. In addition. prior to governmental control. Western culture’s Santa’s team. where the shepherd eliminates the unﬁt and dogs guard their herd from attacks by wolves. Mammoth bio-mass presumably could have been equaled by tasty rodents: lemmings. etc. the fate of dogs and humans has been intertwined for a very long time indeed. however. Unfortunately. voles. even long before the time the hypothetical children of African Eve started to spread into Eurasia. taxation. and this process is still going on in our time. apparently having some success in killing mammoths. Schleidt/Michael D. Reconsideration of past and current concepts of domestication has become inescapable. that those wolves never became specialized big game hunters like the large cats of Africa. Shalter There is also indication of later inﬂuxes of wolf genes into dog populations. Well. Very much like today. From a biologist’s vantage point. Wolf: The Pastoralist In fact. so it has been proposed that Neanderthals similar to present-day gatherer-hunters also collected other foodstuffs (fruits. the reindeer. and at times continued into the continent of North America? Although the mammoth must have been an impressive sight. and to cooperate. very likely. the sick and the aged. human pastoralists of reindeer. No. and hyenas. weaklings. what about the state of wolfkind when Neanderthals roamed the “Mammoth Steppe”. and special funding. bears. Wolf packs. It is noteworthy. however. and before bounty hunting and fur trade. in shepherd-controlled sheep herds. wolves could have played a similar pastoral role: eliminating the unﬁt and keeping away the big cats.). to share risk fairly among pack members. whereas wolves take only what herd owners would never touch and would not even feed their own dogs: placentas on calving grounds. before it became a national sport to shoot wolves from helicopters. 1 . the Eurasian tundra and grass steppe ecosystems which reached from Spain to the far east of Siberia. by far. it was not the species that constituted the greatest amount of biomass in that ecosystem. there are many more artifacts attributed to Neanderthals than bodily remains. Latin for den11) as suggested earlier (SCHLEIDT 1998) and wonder who cubilicated whom. Vol. the intertwining process of hominization and canization makes sense only if viewed as coevolution. and by the most abundant species of wolf food. It appears to us especially signiﬁcant that the ﬁrst dogs separated from wolves in an area and at a time when Neanderthals were the only hominids within the distribution range of wolves. they may have raided bee hives and underground grain stores of hamsters and other rodents. were an important part of that ecosystem. Neanderthals were wandering around Europe. Wolves ability to hunt as packs. Every new ﬁnd seems to raise more questions than it answers. and occasional images provided by National Geographic are all that is left of a species that was once part of one of the largest ecosystems on earth. not only tolerated wolf packs following their herds. Thus. unsurpassed by any of the big cats. in the ancient form of reindeer management. in Siberia. Wolves retained their full reper- ❘ 66 ❘ 2003. So. since the meeting of wolves and modern humans predates. basically living off the herds they followed in their annual cycle of migration. we should talk about “cubilication” (cubile. etc. we may ask: What was the state of affairs among our ancestors when some wolves separated from their conspeciﬁcs and became the immediate precursors of dogs? At that time. has not been integrated into the descent of modern hominids. Hominization and Canization We now are confronted with this startling temporal and geographical coincidence between the emergence of mankind and dogkind. Consequently. our primate heritage. and much of what is being taught one day by one school of thought is chalEvolution and Cognition lenged by another the next. but even considered those wolves to contribute to the breeding of better reindeer. between hominization and canization. Consequently. Few animals can live on meat alone. has attracted attention ever since the publication of Darwin’s The Descent of Man. Nowadays. Canids’ use of dens dates back further. The descent of dogs.Wolfgang M. squirrels. to the best of our knowledge. instead of domestication. Thus. ahead of human pastoralists by tens of thousands. Eurasian wolves can be viewed as the ﬁrst true pastoralists. Even the term domestication has the wrong ring. Humans select only the best specimens for slaughter. Whereas the evolution of man. hundreds of thousands of years. among mammals. a few specimens in various zoos. 9. greens. the evolution of wolves and dogs has remained a particular topic for paleontologists specialized in Pleistocene carnivores. moved them to the top of the food pyramid on the Eurasian plains. horses. anything that could be considered a human habitation in the form of a domus (Latin for house ).
let’s look at other alternative uses. tending toward a degree of omnivorousness to which many human steak fans would pale in comparison.”(ZEUNER 1963). seasonally in vast herds in the realm between what is now Spain and eastern Siberia. even supplanting the wolves at the top of the food pyramid? A single Neanderthal. however. ZEUNER. and toys or playmates for human children (GROVES 1999). Reindeer. In a fair comparison.g. convincing them that attempts to obtain human meat were not worth the risk (also still the best argument against human cannibalism as a stable strategy). having made it successfully through several ice age climate changes. at least since around 400 ka PB. but the hunter with his gun will surely get him. they take their aphids out to pasture in the trees. it tends not to run from a small predator. There. Pups could have been used as baby substitutes. in the spring. Adult wolves. As a group. at ﬁrst only as junior partners. and Bushman fooled African undulates using similar tricks. they repay the ants service with aphid honey. at this moment. A single wolf may occaEvolution and Cognition sionally gulp down a little Red Ridinghood. as for example aphids and ants. 6) stakes out the time–space continuum the past 60 Ma BP within three continents: North America. however. or the wolf loses its skin. Africa. which tend not to run away from every little disturbance. we lack evidence for the use of tamed wolves as hunting companions. the aphids thrive and multiply under the close supervision and even protection of their ants. symbiotic. and. and at times crossing the Bering strait into the North American continent. we can assume that early Eurasian hominids. over hundreds of thousands. Even hiking on the Alpine high pastures with a dog can easily provoke such a mobbing response in cattle. Neanderthals were superior to wolves only in (1) having greater cognitive ability and foresight (reﬂected especially in their scouting and scavenging skills). however. had the same mutual relationship with wolves as exists today in comparable cultures and situations. reasoning about the domestication of dogs in his famous 1963 History of Domesticated Animal. The latter is especially signiﬁcant in dealing with herds of ungulates. Vol. Some ants keep aphids throughout the winter in their nests. apparently not to provoke pursuit. (2) seeing better at longer distances (having an eye level twice that of wolves. and (3) being able to hit a distant target. when the ﬁrst leaves appear. and have shared the plenty of those large reindeer herds without raising the level of intra-pack social friction. So. presumably mainly by scavenging and occasionally killing big game. E. after a careful comparison between herding behavior of wolves to single out potential prey and the herding behavior of sheep dogs. hot water bottles. traveling in ancient times. and it is hard to imagine that a man would take the risk of becoming outranked by his former companion much less his former servant. and in addition berries of various kinds. but approach a serious predator with curiosity: American Indians pulled over a wolves hide to get close to a herd of buffalos .12 How could all this have happened around 135 ka BP? The ﬂow diagram at the very end of this paper (Fig. stated that “ the wolf and the pastoralists might be seen to have much in common. 1 . spear and stone weapons and with all his strength.Co-evolution of Humans and Canids toire and zeal for hunting small rodents. were probably a risky addition to a Neanderthal family. it becomes an easy target for a skilled hunter’s spear. could well have coevolved with wolves in the sense that prey and predator became interdependent. e. A single Neanderthal out alone would have been no match against his own wolf’s jaws in a sudden ﬂare-up dispute over rank. Homo erectus. Man: The Reindeer Hunter Could Neanderthals around 135 ka BP. in the end. taking birds and eggs. what would Neanderthals and wolves have gained from a cooperative coalition? Since. and. maybe millions of years. F. Thus. a group of Neanderthals could have eased their way into the thriving business of wolf pastoralists. No. even armed with ﬁre.. wound and even kill several members of the pack. in exchange. Neanderthals undoubtedly could keep a pack of wolves at bay. literally cut away by his former servant’s fangs. armed with ﬁre and spears (THIEME 1997). protected by wolves? Had Neanderthals already ganged up with canids. Standing still. Another common behavior among herding ungulates is standing one’s ground: When a single individual has been separated from its herd. also have entered into the hypothetical ecosystem of the large herds of reindeer. 9. the bond between such a brave hunter and his wolf would have been broken: either the hunter loses his face. would have been no match for a pack of wolves out to have him for a meal. So. Thus. and Eurasia as centerﬁeld where “Man ❘ 67 ❘ 2003. in the ensuing struggle. able to cover four times an area in the steppe).
due to new insights into the effect of continental drift on ocean currents and air streams. but a mixture of “the best of all our ancestors”. more successful ﬁshermen. better hunters. presumably followed by the aurochs ❘ 68 ❘ 2003. Wolfkind Today Once a few Neanderthals had learned to live with wolves and adopt the pack algorithm (going beyond the close ties of kinship. we constructed a basic framework for the coevolution of hominids and canids: depicting the global time–space continuum on a logarithmic time scale. technology transfer and diversiﬁcation began to thrive. only faster. and of all the non-human mammalian species that roamed Eurasia 1 Ma BP. Vol. There are many more variables of fundamental importance for the evolutionary process: food plants: e. One more point about climate is commonly not even mentioned. and cores from drilling deep sea bottoms and ice caps. beasts of burden.g. infectious diseases. Today.. not shown in this schema for reasons of clarity. there was already some ﬂint trade going on across Europe. and the interactions of even the most relevant biological variables in that complex ecosystem and its effect on evolution. wolves were the most successful in increasing their numbers as dogs. What has been named “Holocene”— our current epoch in which humanity has begun to change the face of the earth—is just an interlude before the next cold phase (Fig. namely that the remarkable Pleistocene ﬂuctuations did not roast and chill our ancestors and their dogs in a 100 000 year rhythm. climate. Schleidt/Michael D. that is. Especially our understanding of the climate of the past has dramatically changed during recent years. the winters. from the smallest plants and insects to the largest mammals. Glaciers had receded around the middle of the Riss glaciation. bordered by rolling blizzard swept steppe and glaciers is still deeply engrained in teaching and described in textbooks. but in summer the temperature in the valleys and in the plains rose nearly as high as at present and supported vegetation similar to that of Central Europe today. guards. coding basic climatic conditions and indicating the conﬁnes of the continental plates and narrow land bridges. baby substitutes. But. should we think that all wolves got attached to all reindeer. and vegetation. insect pests. by a more detailed. Another important aspect is that. and sharing risks) many alternative ways to make a living became available.Wolfgang M. and along such routes. Humans became better gatherers. 1 . Considering for how long Homo erectus type hominids were residents of Eurasia as far as China and for how long Neanderthals did very well for an amazingly long time. subdivided by brief interstadial and interglacial warm spells. or moved with humans. human substitutes in experiments. sled pulls. ﬁne-grain analysis of local geography. the temperatures Evolution and Cognition were much higher than today. wherever they were. As the trees. in the interglacials. where ultimately everything is connected to everything. 5). All this must have resulted in major population changes. the image of the ice age landscape as endless plains of inhospitable snow-covered permafrost. learning to cooperate closely. etc. Within this process of coevolution. because they move with legs and not by seeds and their selective survival. gardeners. tame wolves could have changed hands. We personally favor the idea that we are not the descendents of an African elite. and after another advance of the glaciers and loess steppe. lichens and grasses. Reindeer are mostly out of sight. and that all Neanderthals lived off mammoths and cave bears at ﬁrst and then suddenly switched to reindeer to become helpers to wolves. Unfortunately. deciduous trees thrived in Europe’s Riss-Würm interglacial period. Thus. 9. as human genes and customs spread from one group to the next. food. you name it. by no stretch of the imagination. Shalter meets dog” in a coevolutionary process.. a more realistic picture of that important time span of human history. and areas now famous as European ski resorts supported Mediterranean vegetation. Yes. at least. man sits atop the food pyramid throughout the entire world. “Pleistocene” is not simply a past epoch of grueling freezes. and new records of the past climates from pollen analysis. as it best suits our needs to understand the weave of nature. However. so did the animals. And. Wolves became hunting companions. toys. “we are” in Pleistocene. No. Only slowly do we gain. predators: notably the big cats. but moved gradually to stay within their preferred ranges of temperature and humidity. grasses and ﬂowering plants did not die out. astronauts. Within this framework species can be added or deleted. especially in areas covered by glaciers were long and hard. coniferous trees had reappeared. and the ﬁrst “astronauts” to circle our planet. We are still far from understanding all the consequences of the dramatic changes of climate. rather. it is hard to believe that they disappeared without leaving a trace within the human genome.
When grassland opened and herds of grazing ungulates. BUDIANSKY’s idea that wolves weaseled their way into our hearts as scavengers). originally about the size of a fox. but rather a proposal of an alternative that we ﬁnd equally reasonable. In the center an estimate of global climate changes during the past 1. In fact. 1 . Wolves meeting humans in a phase of the latter’s apprenticeship in wolf pastoralism and. Africa. and the Asian dhole—became part of the rich palette of predators and scavengers. based on the familiar algorithm: “The sun will certainly rise tomorrow because it always did”. sapiens “EVE” Homo s. coex- now represented by our cattle.g. raping. wolves conquered Africa (e. effecting temperatures. no doubt. sapiens “ADAM” Hopmo s.Co-evolution of Humans and Canids Homo sapiens Homo s. ocean level. intercontinental faunal exchange (the extent of polar ice caps. We could not resist extrapolating to the future glaciation (indicated by “???”). based on oxygen isotope ratios in cores samples from the Ocean Drilling Program. ensure justice. coyotes (all Members of the genus Canis). Homo homini Pithecus— Lupus homini Homo? Figure 5: Climate and evolution of canids and hominids. jackals. notably horses and antelopes began to dominate the open plains of North America.g. estimates based on V ILÀ et al. or killing. there is some overlap. and around 10 Ma BP. and subsequently. and Antarctica. Homo homini lupus has been cited whenever humans turn on each other. plundering. is a good alternative hypothesis to the current theories of domestication with man conquering beasts. they started to cross into Asia.(right tree. some early canids moved into this new habitat. and back into North America. with a leaning toward omnivorousness. Less global ice ??? 0 Würm Riss More global ice ka BP 200 400 600 800 1000 1200 3 4 5 δ18O(‰) Mindel Günz Dogs Dogs & Wolves Dogs Wolves Wolves Dogs & Wolves Coyotes Are Wolves That Beastly? The human is like a monkey to other humans. akin to the feeding habits of the bears. and the aberrant “wild dogs”: the African Cape hunting dog. 9. We advocate the exploration of this alternative in the belief that within the complex process of co-evolution at different times and localities. The labels Würm. Evolution and Cognition ❘ 69 ❘ 2003. antisocial and selﬁsh led Thomas HOBBES (1588–1679) to recommend. A multitude of swift canid predators co-evolved with the herding ungulates. and have not ended today. Thus. This pessimistic view that human nature is essentially brutish. 1997) and of hominids. No. closer to the biological evidence. based on various current sources). a variety of interactions between humans and canids could have occurred that shaped their future interdependence. or by sheep. Mindel and Gunz refer to the four major Alpine glaciations. wolves becoming dogs and early humans becoming modern man. Or. Final Remarks This is not a POPPERIAN attempt to falsify any speciﬁc hypothesis concerning the evolution of humans or canids. Polynesia. the canids—the wolves. an all-powerful central government to impose order. in a subsequent process of coevolution. Is the wolf like a human to other humans? Well. in his treatise Leviathan. van H USEN 2000) Evolution of canids (left tree. robbing. The canids also known as (wild) “dogs” have their roots in North America. and prevent men from destroying themselves in bestial fashion (HOBBES 1985). through cognitive superiority and to the bootstrapping theory of hominization with man domesticating himself (e.2 million years. Early accounts of our forefathers brutish behavior in the Bible and stories throughout history describe such atrocities.. as the Basenji infringed on the Lycaon range) and used humans as a vector to get to Australia (dingo). including wolves. attacking for no defendable reason. They were forest dwelling carnivores. Riss. Vol. their closest relatives. Europe. and some dogs behave more humanely than some humans. neanderthaliensis Homo erectus Australopithecus I Vilà 97 II Vilà 97 III Vilà 97 IV Vilà 97 Homo Homini Lupus? “Man to Man is an arrant Wolfe” (HOBBES 1651). where Würm corresponds to the Northwest-European Weichsel and to the North-American Wisconsin glaciation.
The advantage the canids had over their competitors was their special ability to deal with the herdrecent 1. vive by scavenging. and neither As noted above. as seeing a big stick and speaking softly (at least. move fast and enduringly. early humans turned into omnivorous complemented human skills and satisﬁed human gatherers and scavengers. as a species distinct from the rent state of the evidence concerning canid and human cocoyote. Michael mild interglacials of the Ice Wolfgang M. hunter–gardeners (Homo erectus) into the open savanna. by carryservice. Age. As cunning scavenAcknowledgments gers. and at times of need their ability to surstyle as herd followers and herders of reindeer. Thus. and Kurt KOTRSCHAL. hot-water-bottles. Today’s “wolf-type canids” are Tertiary 10. 9. Wolves and huthe social canids remained the dominant predator. cape hunting dog and dhole.schleidt@univie. bears Africa Eurasia North America and mustelids. exploiting an ecological niche that anticdence occurred: at some time during the last ice age.000 ing strategies of the ungulates. In the absence and. around 150 ka BP the Email: retlash@aol. At this point. the “wolf-type Riss 100.000. diaper multitude of resources.. and her daughters entered the Neready several Ma BP. MeanMichael D. and make most efﬁcient use of Mindel Günz every single kill by their ability to “wolf down” a 1. ipated early forms of pastoralism. too numerous to menand adjoining Asia. appeared 1 Ma BP. culminating in the sucEmail: wolfgang. they moved into the Authors’ address Special thanks especially to plains of Eurasia during the Dorothy W. canids became herd anderthal domain. With this wide range of abilities horses. a strange coincifollowers. at ﬁrst) ing-eye dogs. Schleidt W. e. thus nipping Würm away stragglers and individuals on the margins of a “selﬁsh herd” (HAMILTON 1971). became agriculturists. the cirFigure 6: Graphic summary of the authors’ reading of the curcumpolar big “wolf”. a trait that dates back to around 5 Ma BP. some humans adopted the wolves’ life small prey. Schleidt/Michael D. No. ultimately. While the big cats had become specialized for stalking and surprising 10. And dogs of fruit trees. In the fringes of their range. who helped us in develwhile.000. big cats’ position on the very top of the food pyramid—with the lion as the “king” in the animal kingdom—had been relinquished to the social canids alEve had emerged. not all wolves had become pastoralists.000 large part of the quarry before the scavengers had detected the kill. Shalter tion.000 Pleistocene canids” were able to keep pace with the herds. First.000 considerably more social than any other predators.org/wolfgang/ friends. and Canis lupus. their skills for hunting ure 6). GRACEY.ac.Wolfgang M. 1 . e. hyenas. humankind separated from chimhad all humans. In fact. it appears that the evolution. hunter–gatherers. they learned to discriminate among a beasts-of-burden. Shalter isting and competing with the cats. mans had found their match. and as true trusted companions.at but also to all the other cessful Neanderthal of Europe http://www. or specialized as in Africa around 6 Ma BP and moved as true humans ﬁsh–hunters. they generously share their loot with other pack members. The cast With fruit trees of human and reindeer to the common ancestor of the three social canids: and canine as parts of evolution the stage set wolf. And they never our ancestors teamed up with pastoralist wolves (Figlost their omnivorous habits. to avoid peril. and other hoofed animals. FOX. Thanks to their superior needs in many ways beyond herding and hunting: as brain power.g. and to bluff the ﬁerce predators into deserting their quarry. tribe of the legendary African Evolution and Cognition ❘ 70 ❘ Years before present 2003. forceful attack. The Genus Canis apparently evolved in Asia. kind to top of the food pyramid.com oping our ideas. and “dogs” diversiﬁed until the invention of ﬁrearms propelled humanand moved into other human cultures. Of course. Vol.000 their prey in a sudden. guards.schleidt.g. humans panzee-like tree-dwelling and fruit-eating ancestors remained gatherers and scavengers.
D./Woldegabriel. L. M. Of course. tamed wolves and dogs also have the potential to be subject to abuse by humans (e. CA Fox. (1997) Bonobo: The Forgotten Ape. Van Nostrand Reinhold Company: New York. Clark. http://www. 2003) only supports a presence in that region at that time. F. W. Behavioral Biology and Evolution.C. Journal of Theoretical Biology 31: 295–311.g. De Waal. Vienna on 25 September 1997. R. Royston: London. and we can well assume that in addition to pelage and skeleton behavioral traits vary as well. we take the liberty of borrowing for our brief digest of human evolution a few of those popular terms. (eds) (2003) Wolves: Behavior. F. (eds) (1998) Evolution of Tertiary Mammals of North America.. R. U. D. 11 Same root as in cubicle and concubine. In: Ulrich. S. C. R. and frequently changing emphasis. K. T. (1937) The management of sledge dogs. Ecology. and one may wonder whether the “genus” Canis should be considered a single species. P. L. 4 Jane GOODALL im Interview: „Ich hatte schon während meiner Kindheit einen wundervollen Lehrer: meinen Hund Rusty. M. Janis. (ed) (1975) The Wild Canids: Their Systematics. W. R.. (1982) Early European Agriculture. M. D. Washington. D. but also the benefit of being nursed back to full strength. Harrington. Brown: Boston. R. For example.-R. A. University of Chicago Press: Chicago. P.g./Boitani. (1954) Man Meets Dog. T. L. (2000) The Truth about Dogs. 12 The association between humans and reindeer in Pleistocene Europe had early on led to speculation that reindeer herding is an ancient form of subsistence (J ARMAN/BAILEY/ JARMAN 1982).1518. Originally published in 1651. W. Vienna). N. J. (1985) Leviathan. in our considered opinion. L. we restrict our review to a few essentials and abstain from citing specific references for specific statements.und Spätpaläolithikum. 3 The paleontological record of canid evolution is quite rich.. (1980) The Soul of the Wolf. H. Budiansky. Forme. S. 6 Of course. Jarman./Boisserie. or the Matter. P UCHER 1986). Our review is based mainly on JANIS/SCOTT/JACOBS (1998) and MUNTHE (1998). WOLPOFF/CASPARI 1997)./Renne. C. E. In the other extreme. dass wir Tiere sind”. Macpherson./Hart./Gilbert. Methuen: London.g. B. Volume 1. (Archive Wolfgang M. (eds). & Power of a Common-wealth Ecclesiasticall and Civill. Ethiopia. Viking: New York./Suwa. Henry Holt: New York./Kathoh. the readings of the human record are still highly diverse and personal. Cambridge Univ. Mech. L. Hobbes. (1971) Geometry for the selfish herd. (ed) Man and Environment in the Palaeolithic. (eds) (1982) Wolves of the World./Bell. Lorenz. W. chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash. Mech. G./Bird./Jacobs./Paquet.“ (BAUR 2001)./Scott. 2 Following the common practice of balancing the paucity of the archeological record by generous interpretations in plain language. Evolution and Cognition ❘ 71 ❘ 2003. C. J. (1997) Fax to Wolfgang M. further details in M UNTHE (1989). Vol. (1999) The advantages and disadvantages of being domesticated. who knew of multiple matings of the queen all along. G./Asfaw. D. M. Edited with an introduction by C. (2001) Jane Goodall im Interview: “Von den Schimpansen lernen.de/wissenschaft/erde/ 0. considering the scarcity of early Homo fossils (compared to the fossil record of Canis). K.I. and Conservation. Columbia University Press: New York. University of California Press: Berkeley. Guthrie. J. Z. N. Perspectives in Human Biology 4: 1–12. M. and sometimes highly speculative nature of the interpretation of the paleontological record of human evolution. exaggerated claims. T. References Baur. (1970) The wolf: The ecology and behavior of an endangered species. Belknap: Cambridge MA. K. Cambridge University Press: Cambridge UK. Goodall. A.) 215320. (1995) Mensch–Tier Beziehungen im Jung. 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